Pinus nigra - Euforgen

EUFORGEN 

Technical guidelines for genetic conservation and use 

European black pine 

Pinus nigra 

V. Isajev 1 , B. Fady 2 , H. Semerci 3 and V. Andonovski 4 

1 

Forestry Faculty of Belgrade University, Belgrade, Serbia and 

Montenegro 

2 

INRA, Mediterranean Forest Research Unit, Avignon, France 

3 

Forest Tree Seeds&Tree Breeding Research Directorate, Ankara,Turkey 

4 

Faculty of Forestry, Skopje, Macedonia FYR 

These Technical Guidelines are intended to assist those who cherish the valuable European 

black pine genepool and its inheritance, through conserving valuable seed sources or use in 

practical forestry. The focus is on conserving the genetic diversity of the species at the European 

scale. The recommendations provided in this module should be regarded as a commonly agreed 

basis to be complemented and further developed in local, national or regional conditions. The 

Guidelines are based on the available knowledge of the species and on widely accepted 

methods for the conservation of forest genetic resources. 

Biology and ecology 

The European black pine (Pinus 

nigra Arnold) grows up to 30 

(rarely 40–50) m tall, with a trunk 

that is usually straight. The bark 

is light grey to dark grey-brown, 

deeply furrowed longitudinally on 

older trees. The crown is broadly 

conical on young trees, umbrella-shaped 

on older trees, 

especially in shallow soil 

on rocky terrain. 

Branch tips are 

slightly ascending 

on young trees; on 

older trees only 

branches at the top 

part of the crown 

have upturned tips. 

Needles are rather 

stiff, 8–16 cm long, 

1–2 mm in diameter, 

straight or curved, finely 

serrated. Resin ducts are 

median. Leaf sheath is persistent, 

10–12 mm long. 

Black pine is a monoecious 

wind-pollinated conifer, and its 

seeds are wind dispersed. 

Flowering occurs every year, 

although seed yield is abundant 

only every 2–4 years. Trees reach 

sexual maturity at 15–20 years in 

their natural habitat. Flowers 

appear in May. Female inflorescences 

are reddish, and male 

catkins are yellow. Fecundation 

occurs 13 months after pollination. 

Cones are sessile and horizontally 

spreading, 4–8 cm long, 

2–4 cm wide, yellow-brown or 

light yellow and glossy. They 

ripen from September to October 

of the second year, and open 

in the third year after pollination. 

Cones contain 30–40 seeds, of 

which half can germinate. Seeds 

are grey, 5–7 mm long, with a 

wing 19–26 mm long. Germination 

can occur without stratification 

although this technique is 

often used in forest nurseries 

(30–60 day moist +5°C treatment). 

Black pine is an obligate 

seeder under natural conditions. 

Most black pine subspecies 

(see Distribution) grow in a 

Mediterranean-type climate, 

except P.n. nigra which is more 

typically temperate. Bioclimatic 

conditions range from humid 

(800–1000 mm annual rainfall) as

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ropean black pinePinus nigraEuropean black pinePinus nigraEuropean black pineP 

in P.n. mauretanica or P.n. laricio, 

to subhumid (600–800 mm) as in 

P.n. pallasiana in Cyprus, to 

semi-arid (400–600 mm) as 

in P.n. pallasiana in Anatolia. 

The optimal altitudinal 

range of black pine is 

between 800 to 1500 m. 

However, a considerable altitudinal 

variation can be 

observed: from 350 to 1000 m 

in Italy (P.n. nigra) and on the 

Croatian coast (P.n. dalmatica), 

from 500 to 900 m in the French 

Pyrenees and 1600 to 2000 m in 

Spain (P.n. salzmannii), from 

1000 to 1600 m in Corsica (P.n. 

laricio), from 1000 to 2200 m in 

the Taurus mountains and 1400 

to 1800 m in Cyprus (P.n. pallasiana) 

and from 1600 to 1800 m 

in North Africa (P.n. mauretanica). 

Black pine can grow on a 

variety of substrates: limestone 

(e.g. P.n. mauretanica, P.n. dalmatica, 

P.n. pallasiana in Central 

Greece), dolomites (e.g. P.n. 

nigra in northern Italy and Austria, 

P.n. salzmanni in the 

Cévennes, France), acidic soils 

(P. n. laricio, P.n. pallasiana in 

Anatolia, P. n. salzmanni in the 

French Pyrenees) or volcanic 

soils (P.n. laricio in Sicily). 

Black pine is a light-demanding 

species, intolerant of shade 

but resistant to wind and 

drought. It grows in pure stands 

or more rarely in association with 

other pines such as P. sylvestris 

or P. uncinata. 

Distribution 

Black pine extends over more 

than 3.5 million hectares from 

western North Africa through 

southern Europe to Asia 

Minor. Owing to this large 

albeit discontinuous range 

and its large genetic and 

phenotypic variability, it 

is regarded as a collective 

species. Although 

there is no definite consensus 

on its taxonomy, 

six main subspecies can 

be recognized from North Africa 

to the Crimea. 

Pinus nigra mauretanica 

(Maire et Peyerimh.) Heywood 

covers only a few hectares in the 

Rif mountains of Morocco and the 

Djurdjura mountains of Algeria. 

Pinus nigra salzmannii (Dunal) 

Franco (syn: P. n. clusiana, P. n. 

pyrenaica) covers extensive 

areas in Spain (over 350 000 ha 

from Andalucia to Catalunia and 

on the southern slopes of the 

Pyrenees) and is found in a few 

isolated populations in the Pyrenees 

and Cévennes in France. 

It is sometimes referred to as the 

Pyrenean pine. 

Pinus nigra laricio (Poiret) is 

found in Corsica (Corsican pine) 

over 22 000 ha, in Calabria 

(where it is also recognized as P. 

n. l. calabrica, the Calabrian pine) 

and in Sicily. 

Pinus nigra nigra (syn: P.n. 

austriaca Höss, P.n. nigricans 

Host, the Austrian pine) is found 

from Italy in the Apennines to 

northern Greece through the 

Julian Alps and the Balkan 

mountains, covering more than 

800 000 ha. 

Pinus nigra dalmatica (Vis.) 

Franco, the Dalmatian pine, is 

found on a few islands off the 

coast of Croatia and on the 

southern slopes of the Dinaric 

Alps. 

Pinus nigra pallasiana (Lamb.) 

Holmboe covers extensive areas, 

mostly in Greece and Turkey (2.5 

million ha, 8% of total forest 

area) and possibly as far west as 

Bulgaria. It can also be found in 

Cyprus and the Crimea. It is 

sometimes referred to as the 

Crimean pine.

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Importance and use 

Black pine is one of the most 

economically important native 

conifers in southern Europe. Early 

growth is rather fast. It is widely 

planted outside its natural 

range. Wood is durable and rich 

in resin, easy to process. 

P.n. laricio is appreciated 

for building and 

roofing because of its 

straightness and thin branches. 

If properly thinned, its low 

amount of duramen makes it a 

fine carpentry and cabinetry 

wood. The same use can be 

made of Calabrian pine, although 

it is more branchy. Wood of P.n. 

nigra is of lower quality and thus 

restricted to lower-grade building 

wood and the making of crates. 

Black pine has a mean productivity 

of 8–20 m 3 ha -1 yr -1 when 

grown as a monoculture on 

fertile soils. In natural 

conditions, productivity 

is 6–10 m 3 ha -1 yr -1 and 

down to less then 3 m 3 ha -1 

yr -1 on the driest sites. 

Because of its ability to 

develop well on open lands 

and in ecologically demanding 

situations, Austrian pine was 

intensively used during 19 th and 

early 20 th century reforestation 

programmes, e.g. in the French 

southern Alps for landslide control 

and land rehabilitation and in 

England and the USA for sanddune 

fixation and as a windbreak. 

Currently P.n. laricio is the 

most important reforestation 

species in southern England as 

well as in some French regions 

(e.g. Loire valley). 

Black pine is also valued for 

landscaping, both in parks (isolated 

trees or in groups) and in 

urban and industrial contexts 

because of its tolerance to pollution. 

It is one of the most 

common introduced ornamentals 

in the USA. Other uses 

include Christmas trees, fuel 

wood and poles. 

Black pine is included in 

the European Council Directive 

1999/105/CE (December 

22 1999) on the marketing 

of forest reproductive 

material. Minimum 

requirements have to 

be met before black 

pine seed can be 

sold for reforestation. 

Genetic knowledge 

The first black-pine-type fossils 

date to the Miocene, about 20 

million years ago. The ice cycles 

that shaped the Quaternary period 

in Europe are believed to have 

been responsible for the currently 

very discontinuous range of black 

pine. This geographic separation 

did not result in mating barriers, 

and all subspecies are interfertile 

under experimental conditions. 

Studies using morphological and 

genetic markers have confirmed 

the common phylogenetic origin 

of all black pines. The most divergent 

and genetically original European 

groups are P.n. salzmanii 

and P. n. laricio, although P.n. 

nigra, dalmatica and pallasiana 

appear quite similar. The amount 

of genetic diversity is also high 

within populations. Experiments 

measuring adaptive traits have 

revealed strong within- and 

among-population variability for 

traits such as vigour, form and 

drought, frost and disease resistance. 

It is this huge adaptive plasticity 

that has made black pine 

such a favourite for reforestation 

projects over a wide range of 

environments. 

In the middle of the 20 th century, 

several provenance trials 

were established independently 

in Europe, the USA and New 

Zealand. The Corsican and Calabrian 

black pine provenances 

were found to be the best in 

almost every respect on siliceous 

soils. They had consistently

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excellent stem form and branching 

habit, gave the greatest volume 

production and were hardy 

against winter and late frosts 

(except in north-central USA). 

The major defect reported is 

branch forking, which is both 

heritable and highly correlated 

with polycyclism and branch 

angle. On calcareous soils, P.n. 

laricio does not perform well and 

is to be replaced by the slowergrowing 

but more Ca-tolerant 

P.n. nigra. In dry climates (as in 

inner Anatolia, Turkey), black 

pine is slow growing and breeding 

programmes for such zones 

are focused on improving growth 

rate and increasing drought and 

frost tolerance through withinpopulation 

selection. 

Intraspecific hybridization is 

easily performed among all black 

pine subspecies (a further proof 

of phylogenetic relatedness), but 

has not contributed any outstanding 

genotypes to breeding 

programmes so far. Interspecific 

crosses seem to be possible at a 

low survival rate with P. 

sylvestris. 

Black pine seed orchards 

have been established in several 

European countries, e.g. in France 

there are one Calabrian pine and 

two Corsican pine seed orchards. 

Current experiments in vegetative 

propagation include micropropagation 

of zygotic embryos and 

brachyblasts as well as somatic 

embryogenesis. Propagation by 

grafting has been known since 

1820; the method generally used 

is lateral grafting. 

Threats to 

genetic diversity 

Black pine is not recognized as a 

threatened species although 

some of its sub-Mediterranean 

endemic populations constitute 

priority habitats under the EU 

Natura 2000 directive (Habitat 

Directive n° 92/43/CEE, May 21 

1992). 

Extensive plantations were 

often made across Europe in the 

past two centuries with material 

from unknown and/or very distant 

sources for which no historical 

traces currently exist. This 

has probably resulted in extensive 

mixing of local and imported 

genepools all over the distribution 

area of black pine. 

Damaging insects include 

European black pine shoot moth 

(Rhyacionia buoliana), pine processionary 

caterpillar (Thaumetopoea 

pityocampa), especially 

in warm and dry climates, and tip 

blight (Sphaeropsis sapinea), 

which has been particularly 

active in France and Turkey in 

1990’s. Other pests such as 

Acantholyda hieroglyphica, Diprion 

pini, Pissodes validirostis and 

Monophlebus hellenicus have 

been active in Turkey. Most 

recently, an increase in the 

impact of a needle blight known 

as the ‘red band disease’ (Dothistroma 

septospora) has been 

reported.

Pinu 

European black pinePinus nigraEuropean black pinePinus nigraEuropean black 

Distribution range of European 

black pine 

In areas where black pine is 

widespread and very important 

for forestry, factors such as forest 

fires and illegal cutting cause 

serious damage. In areas where it 

occurs in small isolated populations, 

major risks come from any 

factor that may provoke local 

extinction, either through illegal 

cutting and fires or through 

hybridization (‘genetic pollution’) 

from planted black pines belonging 

to other subspecies. Original 

and rare varieties such as P. nigra 

var. pyramidalis or P. nigra var. 

sheneriana in Turkey are under 

identical threats. 

Guidelines for genetic 

conservation and use 

Because black pine of different 

origins has been extensively 

planted, it is now important to 

identify authochthonous populations. 

This undertaking should be 

carried out at the international 

level. In each country, an inventory 

should be made to define 

the geographical distribution of 

the species, its conservation status, 

threats and potential uses. 

Breeding activities provide valuable 

information by defining 

potential plantation, seed collection 

and transfer zones. In situ 

conservation activities should be 

encouraged separately as seed 

stands and gene conservation 

forests. Those do not serve the 

same goal and should not 

always be identical, especially to 

make the conservation of marginal 

populations possible. An 

international in situ network of 

100–120 stands would seem 

appropriate to represent the natural 

ecological and genetic variability 

of black pines. 

As intraspecific hybridization 

is easy among black pines, exotic 

or improved black pines should 

not be planted in the vicinity of 

autochthonous and naturalized 

stands. This is particularly true for 

localized and fragmented sub-

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EUFORGEN 

These Technical Guidelines were 

produced by members of the 

EUFORGEN Conifers Network. 

The objective of the Network is to 

identify minimum genetic conservation 

requirements in the long 

term in Europe, in order to reduce 

the overall conservation cost and 

to improve the quality of standards 

in each country. 

species such as P.n. laricio, and is of extreme importance for subspecies 

that are threatened, such as P.n. salzmanii in France and P.n. 

mauretanica in North Africa. For these subspecies and other varieties 

of rare occurrence, ex situ conservation is appropriate and urgent. As 

a step in that direction, in 1999 a gene conservation forest was selected 

in Turkey for the rare P. nigra var. pyramidalis. 

Information on the provenance and progeny trials established 

throughout Europe should be entered in a database. This network of 

experimental sites could be used for ex situ conservation of black 

pine. Marginal areas might need to be further sampled to strengthen 

this network and possibly planted as ex situ seed orchards to re-install 

depleted resources. 

Citation: Isajev, V., B. Fady, H. 

Semerci and V. Andonovski. 

2004. EUFORGEN Technical 

Guidelines for genetic conservation 

and use for European black 

pine (Pinus nigra). International 

Plant Genetic Resources Institute, 

Rome, Italy. 6 pages. 

Drawings: Pinus nigra, Claudio 

Giordano. © IPGRI, 2003. 

ISBN 92-9043-659-X 

Selected bibliography 

Lauranson-Broyer, J. and Ph. Lebreton. 1995. Flavonic chemosystematics of 

the specific complex Pinus nigra Arn. Pp. 181-188 in Population genetics 

and genetic conservation of forest trees (P. Baradat, W.T. Adams and G. 

Müller-Starck, eds.). SPB Academic Publishing, Amsterdam. 

Nikolic, D. and N. Tucik. 1983. Isoenzyme variation within and among populations 

of European black pines (Pinus nigra Arnold). Silvae Genetica 32(3- 

4):80-89. 

Quézel, P. and F. Médail. 2003. Ecologie et biogéographie des forêts du bassin 

méditerranéen. Elsevier, Paris. 

Tutin, T.G., V.H. Heywood, N.A. Burgess, D.M. Moore, D.H. Valentine, S.M. 

Walters and D.A. Webb, eds. 1983. Flora Europaea, Vol 1, 2nd edition, pp. 

40-44. Cambridge University Press, Cambridge, UK. 

Vidakovic, M. 1974. Genetics of European black pine (Pinus nigra Arn.). Ann. 

Forest. 6/3 JAZU Zagreb:57-86. 

` 

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